[1] Univ Aberdeen, Inst Biol & Environm Sci, Zool Bldg, Aberdeen AB24 2TZ, Scotland
[2] Univ Aberdeen, Oceanlab, Newburgh AB41 6AA, Aberdeen, Scotland
[3] Newcastle Univ, Sch Marine Sci & Technol, Ridley Bldg, Newcastle Upon Tyne NE1 7RU, Tyne & Wear, England